REVIEW

CORRECTED 20 MARCH 2009; SEE LAST PAGE

opments that would push other naturalists toward
an evolutionary vision during the years he worked

Darwin’s Originality
in isolation. By the late 1850s, the idea of pro-
gressive evolution was widely recognized, and the
positive role of individual competition was being
Peter J. Bowler articulated by thinkers such as Herbert Spencer
(Fig. 1). But key aspects of the Darwinian vision
Charles Darwin’s theory of natural selection has been hailed as one of the most innovative were truly original and would not have occurred
contributions to modern science. When first proposed in 1859, however, it was widely rejected to any other naturalist at the time. Here, Wallace
by his contemporaries, even by those who accepted the general idea of evolution. This article provides a good comparison: He too moved toward
identifies those aspects of Darwin’s work that led him to develop this revolutionary theory, the idea of branching evolution driven by local
including his studies of biogeography and animal breeding, and his recognition of the role played adaptation, but even he did not share Darwin’s
by the struggle for existence. insight that the work of the animal breeders throws
light on the process of natural selection.
he publication of Charles Darwin’s On pre-existing ones in a progressive sequence lead- The theory was both original and disturbing.

T the Origin of Species in 1859 is widely

supposed to have initiated a revolution
both in science and in Western culture. Yet there
ing up to humans (5). But if the general idea of
evolution was not entirely new, Darwin’s vision
of how the process worked certainly was. Al-
It was not just that the idea of natural selection
challenged the belief that the world was designed
by a wise and benevolent God. There was a wider
have been frequent claims that Darwinism was though the theory was eventually paralleled by element of teleology or goal-directedness almost
somehow “in the air” at the time, merely waiting Wallace, Darwin had conceived its basic outline universally accepted at the time. Most thinkers—
for someone to put a few readily available points in the late 1830s, after his return from the voyage including Jean-Baptiste Lamarck and Chambers—
together in the right way [for instance (1)]. The of H.M.S. Beagle. He worked on it in relative took it for granted that the development of life on
fact that Alfred Russel Wallace (Fig. 1) indepen- isolation over the next 20 years, until the arrival of earth represents the unfolding of a coherent plan
dently formulated a theory of natural selection in Wallace’s paper in 1858 precipitated the flurry of aimed at a predetermined goal. (This assumption
1858 is taken as evidence for this position. But activity leading to the publication of the Origin. is still preserved in the very term “evolution”; the
Darwin had created the outlines of the theory Historians have quarried Darwin’s notebooks Latin evolutio refers to the unrolling of a scroll.)
20 years earlier, and there were significant dif- and letters to establish the complex process by The explanatory framework centered on the
CREDIT (LEFT TO RIGHT): STAPLETON COLLECTION/CORBIS; HULTON-DEUTSCH COLLECTION/CORBIS; MICHAEL NICHOLSON/CORBIS

Fig. 1. Charles Darwin, Alfred Russel Wallace, and Herbert Spencer.

ferences between the ways in which he and which he developed his theory (6–9). Darwin theory of natural selection challenged this vision
Wallace formulated their ideas. In this essay, I was a highly creative thinker who synthesized a of nature as an orderly pattern of relations.
argue that Darwin was truly original in his think- number of key insights, some derived from his Darwin’s world view was profoundly differ-
ing, and I support this claim by addressing the scientific work and others from currents circulat- ent because he argued that the adaptation of pop-
related issue of defining just why the theory was ing in his cultural environment. Few would now ulations to their local environment was the sole
so disturbing to his contemporaries. accept the claim that evolution by natural se- cause of transmutation. Many people found it
Darwin was certainly not the first to sug- lection was in the air. Darwin approached the hard to see natural selection as the agent of either
gest the idea of evolution as an alternative to subject in a way that was significantly different divine benevolence or of a rationally structured
the creation of species by God. J. B. Lamarck’s from any of the other efforts being made to ex- cosmic teleology. Selection adapted species to an
theory, published in 1809, had been widely dis- plain the history of life on earth. He had a unique ever-changing environment, and it did so by killing
cussed, although generally rejected (2–4). Robert combination of scientific interests that alerted off useless variations in a ruthless “struggle for
Chambers’s Vestiges of the Natural History of him to topics ignored by other naturalists. He existence.” This did not seem the kind of process
Creation of 1844 sparked a debate over the pos- certainly drew on ideas widely discussed at the that would be instituted by a benevolent God,
sibility that new species were produced from time, but was forced by his scientific interests to especially because its essentially “selfish” nature
use those sources of inspiration in a highly orig- meant that a parasitic way of life was a perfectly
School of Philosophy and Anthropological Studies, The Queen's
inal way. natural adaptive response in some circumstances.
University of Belfast, University Road Belfast, Belfast, Northern To some extent, Darwin may have been More seriously for the idea of cosmic tele-
Ireland, BT7 1NN, UK. E-mail: [email protected] merely “ahead of his time,” anticipating devel- ology, Darwin’s supposition that the production

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REVIEW
of the individual variants in a population was the relations among species. Several proposals his theory to predict an orderly pattern of
essentially undirected ruled out any possibility available in the 1830s deflected attention away relations.
that evolution could be shaped by a predeter- from the model of the branching tree (11). It has been argued that Darwin’s move to a
mined developmental trend. There was no ob- William Sharp Macleay’s quinary or circular more historical viewpoint was inspired by German
vious goal toward which it was aimed, and it system of classification supposed that every romanticism [e.g. (12)], but a more practical
did not produce an orderly pattern of relations genus contained five species that could be ar- incentive was provided by the biogeographical
between species. The accusation that the theory ranged in a circle; each family five genera, and soinsights gained on the Beagle voyage (1831–36).
depended on “random” variation indicated the con- on through the taxonomic hierarchy. Chambers’s The Galapagos species provided the most obvious
cerns of his opponents on this score. As Darwin Vestiges of the Natural History of Creation example of how the relations within a group can
himself made clear, variation was certainly caused depicted evolution in terms of parallel lines ad- be explained by supposing that an original pop-
by something (later identified as genetic muta- vancing through a predetermined sequence of ulation became divided up, in this case by in-
tions), but it was not aimed in any one direction stages within each family, driven by force derived dependent acts of migration to oceanic islands.
and, thus, left adaptive evolution essentially open- from individual development. Here, Darwin followed Lyell in seeing that bio-
ended. He allowed a limited role for variation geography must become a historical
shaped by the organisms’ own activities (the so- science, explaining present distribu-
called Lamarckian effect), but this too permitted tions in terms of past migrations,
multiple vectors of change. Evolution had to be extinctions and (for Darwin but not
depicted as a branching tree in which each act of for Lyell) evolutionary adaptations.
branching was the result of a more or less un- Populations divided by geographical
predictable migration of organisms to a new loca- barriers will develop independently
tion. At the same time, Darwin’s theory undermined as each adapts to its new environ-
the old idea that species were idealized types, ment in its own way, and the pos-
fixed elements in a clearly defined natural order. sibility that barriers can be crossed
Species had to be treated as populations of vary- occasionally allows for the branch-
ing individuals, with no fixed limit on the range ing process of evolution that Darwin
of possible variation. conceived in the late 1830s. It was
by approaching the problem of the
The Tree of Life origin of new species through a study
One innovation at the heart of Darwin’s theory of biogeography that Darwin was
seems so obvious today that it is hard for us to led to construct his model of open-
appreciate just how new and how radical it ended, divergent evolution. Wallace
was at the time. Lamarck had proposed that developed a similar model and tested
there might be natural processes adapting spe- it during his explorations in South
cies to changes in their environment. But Darwin America and the Malay Archipelago
was perhaps the first to realize that if adaptation (modern Indonesia).
to the local environment was the only mechanism Adrian Desmond and James
of evolution, there would be major implications Moore have recently proposed that
for the whole system by which species are clas- Darwin’s hatred of slavery prompted
sified into groups. Darwin’s mentor in geology, his move toward evolutionism (13).
Charles Lyell, had shown how his uniformitarian Because many slaveholders argued
theory would allow the biogeographer to re- that the black race was separately
construct the migrations of species on an ever- created from the white, Darwin
changing earth. Populations could sometimes wanted to show that all races share
become divided by geographical barriers, so that a common ancestry, and he realized
what was once a single species could split into that this claim could be defended
multiple branches adapting to separate environ- by extending the idea throughout
ments (10). Evolution would become a divergent the animal kingdom. As a basis for
process, with some branches splitting over and his thinking, this thesis is sure to
over again, whereas others came to a dead end Fig. 2. Tree of Life, from Darwin’s notebooks (22). generate much controversy, but if
through extinction. accepted it would emphasize the
The image of the tree of life had appeared in These rigidly structured models of taxo- crucial role played by his move toward a model
Darwin’s notebooks of the late 1830s (Fig. 2) nomic relations and evolution made good sense of branching evolution based on geographical
and was proposed independently by Wallace in to anyone embedded in a vision of nature as a diversity.
CREDIT: SYNDICS OF CAMBRIDGE UNIVERSITY LIBRARY

a paper published in 1855. Both realized that it predictable, orderly system governed by a divine This model was so radical that many late
explained why naturalists were able to arrange plan. Such a world view made it difficult to ac- 19th-century evolutionists were unable to accept
species into groups within groups, using descent cept that the history of life on earth might be it in full. Ernst Mayr argued that the theory of
from a common ancestor to explain the under- essentially irregular and unpredictable, dependant common descent was one of Darwin’s greatest
lying similarities. Closely related species have di- on the hazards of migration, isolation, and local achievements, in addition to natural selection
verged recently from a common ancestor, whereas adaptation. Darwin was led toward his alterna- itself (14). So it was, but I think Mayr over-
the ancestry of more distantly related forms must tive model in part because he was more inter- estimated the rapidity with which other natural-
be traced further back down the family tree to ested in adaptation than cosmic teleology, thanks ists were converted to the theory. Many of the
find the common point of origin. to the influence of William Paley’s natural the- non-Darwinian theories of evolution proposed
The idea of common descent now seems so ology. Natural selection replaced divine benev- during the “eclipse of Darwinism” in the late 19th
obvious that we might wonder what alternative olence as an explanation of adaptation. Unlike century were introduced with the aim of subvert-
models could have been proposed to account for Macleay and Chambers, Darwin did not expect ing the implications of the principle of common

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 REVIEW
descent (15). The American neo-Lamarckians scale and that could be investigated directly. interbreeding individuals. Traditionally, species
Edward Drinker Cope and Alpheus Hyatt pro- There was a well-developed network of breeders were treated as idealized types with a fixed es-
posed that the evolution of each group should be by this time, and although their ideas about sence, any variation from the norm being trivial
seen as a series of parallel lines moved through heredity and variation were distinctly pregenet- and impermanent. The breeders knew that they
the same hierarchy of developmental stages, an up- ical (like Darwin’s own), they had a very clear could produce huge changes in structure by ac-
dated version of the idea suggested in Chambers’s appreciation of how they produced changes in cumulating normal variations over a number of
Vestiges. The similarities linking the species in a their artificially small populations. The insight generations. When Darwin linked this informa-
genus were due not to a recent common ancestry, that they worked by selection may have been tion with his conviction that species could change
but to parallel trends independently reaching the important (this is the point of contention among indefinitely over time, he was driven toward a
same stage of development. Like Chambers, they experts studying Darwin’s notebooks), but the new form of species concept in which the pop-
endorsed the recapitulation theory (ontogeny breeders certainly taught him one thing. He ulation becomes paramount. The natural range of
recapitulates phylogeny, in the terminology intro- realized that in a domesticated population there variability becomes part of the species’ character,
duced by Ernst Haeckel) and saw evolution as is always a fund of apparently purposeless and not the result of accidental deviations from a
the addition of preordained stages to ontogeny. undirected variation among individual organisms. fixed norm. This is what Mayr called the transi-
Adaptation was not crucial once the basic char- Although convinced that the degree of variabil- tion from typological thinking to population think-
acter of the group was established, and the linear, ity was artificially enhanced under domestication, ing, and although he may have exaggerated the
orthogenetic evolution of the group might even- Darwin, nevertheless, accepted that there must be extent to which Darwin himself made the con-
tually generate bizarre nonadaptive characters some equivalent variability in every wild popu- ceptual transition, the subsequent development of
as a prelude to extinction—the theory of “racial lation. The analogy with artificial selection then the selection theory brought this implication out
senility.” Darwin could make no sense of the allowed him to depict natural selection as a par- more clearly.
theory proposed by Cope and Hyatt, because he allel process in which a few variant individuals, In the debates that followed the publication
could not imagine an evolutionary process driven in this case with characters useful to the species of On the Origin of Species, the analogy with
by predetermined trends. But the fact that such rather than the human breeder, survive and re- artificial selection continued to play a key role
theories flourished in the late 19th century dem- produce. Those with harmful characters are elim- by forcing even Darwin’s critics to think about
onstrates just how radical the theory of open- inated by the struggle for existence, just as the the problems of heredity and variation in a new
ended, divergent evolution was to the naturalists breeder will not permit any animal to reproduce if way (18). Opponents such as Fleeming Jenkin,
of the time. it does not have the character he wants. It was the who saw selection working on large variations
breeders who taught Darwin that variation is not or “sports of nature,” were, nevertheless, still
Artificial Selection directed toward some preordained goal, allowing working within the framework defined by this
These non-Darwinian models were ultimately him to build on his existing conviction that adapt- analogy. For supporters such as Francis Galton,
marginalized by the synthesis of the selection ive evolution must be an open-ended, branching artificial selection helped to clarify the nature
theory and genetics in the early 20th century. process. of both heredity and selection, paving the way
Genetic mutations seemed to be essentially plu- At the same time, the breeders’ attitude for the revolutionary impact of Mendelian
ralistic and undirected, providing just the source toward variation pushed Darwin toward the genetics. The notion of “hard” heredity was
of “random” variation that Darwin’s mecha- view that the species is just a population of introduced in opposition to the “soft” form of
nism required as its raw material.
This later development high-
lights the importance of another
insight gained by Darwin in the
late 1830s, his decision to inves-
tigate the work of the animal
breeders (Fig. 3) and his recog-
nition that their method of artifi-
cial selection offered a useful way
of understanding how the equiv-
alent natural process operated.
The exact role played by Darwin’s
study of breeding in the formula-
tion of his theory is much debated
by historians (16–17), but there
can be little doubt of how impor-
tant the analogy between artificial
and natural selection became in his
CREDIT: AMERICAN MUSEUM OF NATURAL HISTORY

later thinking. In this case, Darwin

was truly unique, because even
Wallace did not take this step and
dissociated himself from the link
with artificial selection expressed
in Darwin’s later writings.
Darwin turned to the breeders
in search of a clue as to how a
population could be changed—
here at least was a situation where
modifications were actually be-
ing produced on a human time Fig. 3. Pigeons (23).

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REVIEW
inheritance implied by the Lamarckian process. But his occasional use of highly individualistic matter of science being “misused” by social
The undirected nature of variation was clarified language allowed him to be perceived as the commentators, because Darwin’s theorizing
both through the study of large populations by apostle of free enterprise. Much of what later would almost certainly have been different
Galton and through the breeding studies of the became known as “social Darwinism” was, in had he not drawn inspiration from social, as
geneticists. Although it took some time for the fact, Spencerian social Lamarckism expressed well as scientific, influences. We may well
geneticists to accept the situation, their studies in the terminology of struggle popularized by feel uncomfortable with those aspects of his
of mutation ultimately endorsed Darwin’s Darwin. theory today, especially in light of their sub-
claim that the only way the environment could This point is important in the context of the sequent applications to human affairs. But if
affect the population was by selection. Modern charge raised by modern opponents of Darwinism we accept science’s power to upset the tradi-
evolutionary developmental biology has reop- that the theory is responsible for the appearance tional foundations of how we think about the
ened the question of whether variation and of a whole range of unpleasant social policies world, we should also accept its potential to
evolution can be quite as open-ended as Darwin based on struggle. Darwin exploited the idea of interact with moral values.
and his followers believed. But the non- the struggle for existence in a way that was
Darwinian vision of evolution unfolding to an unique until paralleled by Wallace nearly 20 years
References
orderly, predictable plan has been essentially later. Their theory certainly fed into the movements 1. L. Eiseley, Darwin’s Century: Evolution and the Men Who
marginalized by acceptance of the key insights that led toward various kinds of social Darwinism, Discovered It (Doubleday, New York, 1958).
on which Darwin based his theory of natural but it was not the only vehicle for that transition 2. P. Corsi, The Age of Lamarck: Evolutionary Theories in
selection. in the late 19th century. It did, however, highlight France, 1790–1830 (Univ. of California Press, Berkeley,
1988).
the harsher aspects of the consequences of strug-
3. A. Desmond, The Politics of Evolution: Morphology,
The Struggle for Existence gle. The potential implications were drawn out Medicine and Reform in Radical London (Univ. of
One of the most disturbing aspects of Darwin’s even more clearly when Galton argued that it Chicago Press, Chicago, 1989).
theory was its appeal to the struggle for existence would be necessary to apply artificial selection to 4. P. J. Bowler, Evolution: The History of an Idea (Univ. of
as the natural process that equates with the breeder’s the human race in order to prevent “unfit” indi- California Press, Berkeley, ed. 3, 2003).
5. J. A. Secord, Victorian Sensation: The Extraordinary
activity as a selecting agent. This very harsh vision viduals from reproducing and undermining the Publication, Reception, and Secret Authorship of
of nature certainly threatened the traditional belief biological health of the population. This was the Vestiges of the Natural History of Creation (Univ. of
in a benevolent Creator. The term “struggle for eugenics program, and in its most extreme man- Chicago Press, Chicago, 2000).
existence” occurs in Thomas Robert Malthus’s ifestation at the hands of the Nazis, it led not just 6. D. Kohn, Ed., The Darwinian Heritage: A Centennial
Retrospect (Princeton Univ. Press, Princeton, NJ, 1985).
An Essay on the Principle of Population, although to the sterilization but also to the actual elimi- 7. J. Browne, Charles Darwin: Voyaging (Jonathan Cape,
used in the context of tribal groups competing nation of those unfortunates deemed unfit by the London, 1985).
for limited resources. Darwin saw that population state. Did Darwin’s emphasis on the natural elim- 8. C. Darwin, The Power of Place (Jonathan Cape, London,
pressure would lead to competition between in- ination of maladaptive variants help to create a 2002).
9. M. J. S. Hodge, G. Radick, Eds., The Cambridge Companion
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it might represent a means by which the popu- came possible? 10. M. J. S. Hodge, Stud. Hist. Biol. 6, 1 (1982).
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process worked by eliminating the least fit var- itself a creative force in nature, Darwin intro- Early Nineteenth-Century British Biology (Univ. of
Wisconsin Press, Madison, 1983).
iants within the population and allowing the bet- duced a new and profoundly disturbing insight
12. R. J. Richards, The Meaning of Evolution: The
ter adapted to survive and breed. This was what into the world, an insight that seems to have Morphological Construction and Ideological Reconstruction
the philosopher Herbert Spencer would later refer resonated with the thinking of many who did not of Darwin’s Theory (Univ. of Chicago Press, Chicago,
to as the “survival of the fittest.” Strictly speak- understand or accept the details of his theory. 1992).
ing, natural selection requires only differential re- Darwinism was not “responsible” for social 13. A. Desmond, J. R. Moore, Darwin’s Sacred Cause: Race,
Slavery and the Quest for Human Origins (Allen Lane,
production among variants, but Darwin thought Darwinism or eugenics in any simple way. After London, 2009).
that the pressure of competition was necessary to all, some early geneticists endorsed eugenics by 14. E. Mayr, One Long Argument: Charles Darwin and the
make it effective. It seems that without the input analogy with animal breeding even while dis- Genesis of Evolutionary Thought (Harvard Univ. Press,
from Malthus, he would not have come up with missing natural selection as the mechanism of Cambridge, MA, 1991).
15. P. J. Bowler, The Eclipse of Darwinism: Anti-Darwinian
the theory. evolution. And the Nazis wanted to purify a fixed Evolution Theories in the Decades Around 1900 (Johns
The idea of struggle was pervasive in the racial type, which they certainly did not want to Hopkins Univ. Press, Baltimore, MD, 1983).
literature of the period, but could be exploited admit had evolved gradually from an ape 16. R. J. Richards, Stud. Hist. Philos. Sci. 28, 75 (1997).
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18. J. Gayon, Darwinism’s Struggle for Survival: Heredity and
had already seen how competition could be turned primarily through the elimination of useless var- the Hypothesis of Natural Selection (Cambridge Univ.
into a very different, and in some respects less iants, Darwin created an image that could all Press, New York, 1998).
disturbing, mechanism of progress (21). For too easily be exploited by those who wanted 19. P. J. Bowler, J. Hist. Ideas 37, 631 (1976).
Spencer, the interaction between individuals stim- the human race to conform to their own pre- 20. A. Desmond, J. R. Moore, Darwin (Michael Joseph,
London, 1991).
ulated their efforts to adapt to the changing so- existing ideals. In the same way, his populariza-
21. M. Francis, Herbert Spencer and the Invention of Modern
cial and physical environment. He then invoked tion of the struggle metaphor focused attention Life (Acumen, Stocksfield, UK, 2007).
Lamarck’s concept of the “inheritance of ac- onto the individualistic aspects of Spencer’s 22. C. Darwin, Transmutation Notebook D, from Natural
quired characteristics” to explain how these self- philosophy. Selection portfolio (Cambridge Univ. Library,
Modern science recognizes the importance Cambridge, 1838); P. H. Barrett, P. J. Gautrey,
improvements accumulated over many generations,
S. Herbert, D. Kohn, S. Smith, transcribers and Eds.,
leading to biological evolution and social progress. of Darwin’s key insights when used as a way Charles Darwin’s Notebooks (British Museum of
Spencer’s self-improvement model of progress be- of explaining countless otherwise mysterious Natural History, Cornell Univ. Press, Ithaca, NY, 1987);
came immensely popular in the later 19th century, aspects of the natural world. But some of those available at the Darwin Digital Library, http://
and because it too seemed to rely on struggle as the insights came from sources with profoundly darwinlibrary.amnh.org/.
23. G. Neumeister, Das Ganze der Taubenzucht (B. F. Voigt,
motor of change, it was often confused with the disturbing implications, and many historians
Weimar, 1876).
Darwinian mechanism. In fact, Spencer thought now recognize that the theory, in turn, played
that all humans will eventually acquire the faculties into the way those implications were devel-
needed to interact harmoniously with one another. oped by later generations. This is not a simple 10.1126/science.1160332

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 CORRECTIONS & CLARIFICATIONS

ERRATUM Post date 20 March 2009

Reviews: “Darwin’s originality” by P. J. Bowler (9 January, p. 223). On page 226, reference

8 should read as follows: J. Browne, Charles Darwin: The Power of Place (Jonathan Cape,
London, 2002). In reference 22, Transmutation Notebook D should have been Notebook B.
Also in reference 22, two page numbers were missing: Natural Selection, p. 36, and
Charles Darwin’s Notebooks, p. 180.

www.sciencemag.org SCIENCE ERRATUM POST DATE 20 MARCH 2009 1